(Elasmobranchii: Carcharhiniformes) in the Pliocene of the Mediterranean Sea

N. Jb. Geol. Paläont. Abh. 295/2 (2020), 129–139 Article E Stuttgart, February 2020

The extinct catshark Pachyscyllium distans (PROBST, 1879) (Elasmobranchii: Carcharhiniformes) in the Pliocene of the Mediterranean Sea

Alberto Collareta, Marco Merella, Frederik H. Mollen, Simone Casati, and Andrea Di Cencio With 4 ﬁgures and 1 table

Abstract: Sharks assigned to the carcharhiniform family Scyliorhinidae account for about 160 extant species placed in 18 genera. Most living scyliorhinids are small- to medium-sized ground sharks provided with cat- like eyes and nasal barbels similar to whiskers; hence their vernacular name, “catsharks”. Living catsharks mostly inhabit deep or rather deep waters of the warm and temperate seas worldwide, foraging on small fishes and inverterbates. In the present paper, we report on a lateral tooth of Scyliorhinidae collected from a clay pit at Certaldo (central Italy), where marine mudstones belonging to the famously fossiliferous Pliocene successions of Tuscany are exposed. This catshark specimen represents the second bona fide record of the extinct premontreine species Pachyscyllium distans in the Pliocene of the Mediterranean Sea, as well as the geologically youngest confirmed occurrence of this species worldwide. In the Mediterranean Pliocene, P. distans thus coexisted with the similar but distinct species Pachyscyllium dachiardii. After having been widespread in Northern Atlantic, Paratethyan, and Mediterranean waters in Miocene times, P. distans became confined to the Mediterranean Sea during the Pliocene. Therefore, similar to what has recently been suggested for P. dachiardii, we hypothesise that the range of P. distans contracted southward as colder conditions took hold in the Northern Hemisphere. The eventual extinction of P. distans might be related to the first cooling episode that significantly affected the Mediterranean biota around 3 Ma.

Key words: Chondrichthyes, Scyliorhinidae, Premontreinae, Valdelsa Basin, blue clays, Tuscany, climate change, palaeobiogeography, palaeoichthyology.

1. Introduction deep or rather deep waters (Cappetta 2012). Extant catsharks feed mostly on invertebrates and small fish- The carcharhiniform family Scyliorhinidae includes es; however, the life histories of several living scylio- around 160 living species placed in 18 genera (Weig- rhinid species are still very poorly known (Compagno mann 2016; White et al. 2019); as such, it represents 1984). one of the largest living families of sharks. Most ex- For almost two centuries, the Pliocene marine suc- tant scyliorhinids are small- to medium-sized (i.e., less cessions of Tuscany (central Italy) have been the lo- than 1.5 m in total length) ground sharks that exhibit cation of important finds of fossil remains of elasmo- lengthened, cat-like eyes and, sometimes, whisker- branchs (e.g., Lawley 1876), mostly shark teeth and like nasal barbels (Compagno 1984; Cappetta 2012); batoid dental plates and dermal spines (Marsili 2006, hence their vernacular name, “catsharks”. They inhabit and references therein). These fossils were collected seabottoms in tropical and temperate seas all over the to form important palaeontological collections, such world, ranging from very shallow, marginal-marine ar- as the famous “Lawley collection” (Manganelli et al. eas to depths of more than 2000 m (Compagno 1984). 2006), which was studied (in the past) by illustrious That said, most of the Recent scyliorhinids live in naturalists (e.g., Bassani 1901; De Stefano 1909).

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Fig. 1. Location of Certaldo, the locality where the fossil scyliorhinid tooth GAMPS-00924 was found (black star), and schematic regional geological map. B-CI = Baccinello-Cinigiano Basin; CA = Casino Basin; RD = Radicofani Basin; SI = Siena Basin; VC = Val di Chiana Basin; VE = Valdelsa Basin; VO-RA-CH = Volterra-Radicondoli- Chiusdino basins. Mod- ified afterMartini et al. (2013) and Spadini & Manganelli (2015).

Following the emergence of modern palaeontologi- 2. Stratigraphic framework cal research standards, however, these historical collections have lost some of their scientific importance The scyliorhinid fossil tooth described herein was dis- due to widespread uncertainties regarding their geo- covered by one of the authors (S.C.) at an abandoned graphical and stratigraphic context (Landini 1977; quarry at the periphery of the Certaldo village (Flor- Cigala Fulgosi et al. 2009; Collareta et al. 2016). ence Province, Tuscany, central Italy) (Fig. 1). The Now however, the recent reports of rare and elusive GPS geographic coordinates of our find are the fol- elasmobranch taxa from the Pliocene deposits of lowing: 43° 34′ 08″ N; 11° 01′ 51″ E. Tuscany (Spadini & Manganelli 2015; Collareta The fossil specimen was collected from deposits et al. 2017a; Collareta et al. 2017b; Collareta et al. belonging to the Argille Azzurre Formation, a litho- 2018; Manganelli & Spadini 2019), as well as the stratigraphic unit that largely consists of mostly mas- discovery of new fossil localities that are rich in shark sive, shelf mudstones (hence their formational name, and ray teeth (Bianucci et al. 2019), are contributing meaning ‘blue clays’), whose deposition in this area to shed new light on the late Neogene cartilaginous is referred to the Pliocene (Costantini et al. 2004). fish faunas of the Mediterranean Sea. According to Benvenuti et al. (2014), in the Valdelsa In the present paper, on the basis of a single lateral Basin, an intraformational unconformity divides the tooth collected from Pliocene marine mudstones ex- Argille Azzurre Formation into ‘lower’ and ‘upper’ posed at the locality of Certaldo (Tuscany), we provide blue clays, dated to the basalmost Zanclean and upper the second unambiguous record of the extinct catshark Zanclean – lower Piacenzian, respectively. The suc- species Pachyscyllium distans (Scyliorhinidae: Pre- cession exposed at Certaldo belongs to the upper blue montreinae) from the Pliocene of the Mediterranean clays (= ‘upper Argille Azzurre’ of Nalin et al. 2016) Sea and briefly discuss its palaeobiogeographical sig- and can be referred to the S3 Synthem recognised in nificance. the Valdelsa Basin by Benvenuti et al. (2014). The S3

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Fig. 2. The Pliocene deposits exposed at the abandoned Certaldo quarry. A – General view of the outcrop. B – Collection of fossil invertebrate and vertebrate remains at the base of the partly revegetated artificial cliff (white arrow). C – Close- up view on a fossil- rich horizon cropping out a couple of meters below the top of the cliff.

Synthem is a large-scale depositional sequence that sis spp.) have been calibrated to 3.98 and 3.19 Ma, re- consists of a coarse- grained, deepening-upward delta spectively (Violanti 2012). front stratal package, overlain by a fine- grained divi- At the study site, strata belonging to the Argille Az- sion from an open shelf setting (Dominici et al. 2018). zurre Formation crop out along a partly revegetated The latter comprises a rather monotonous mudstone artificial cliff (Fig. 2A). The succession exposed at the succession that, in the study area, features several Os Certaldo quarry is mostly comprised of substantially trea- and Serpulorbis-rich shell beds (Dominici et al. structureless, greyish-bluish mudstones that host com- 2018). mon remains of macro-invertebrates and erode in a According to Benvenuti et al. (2014), the mud- badland- like fashion (Fig. 2B). Fossils concentrate in stones belonging to the S3 Synthem bear planktonic a ca. 25-cm-thick horizon (Fig. 2C) that also feature foraminifer assemblages indicative of the Mediter- rare invertebrate vertical burrows; this interval runs ranean Pliocene (MPL) zone 4 of Cita (1975) and subhorizontally a couple of meters below the top of the Sprovieri (1992), whose bounding bioevents (i.e., the cliff. The invertebrate assemblage detected at the study last common occurrence of Globorotalia margaritae site includes several species of bivalve and gastropod and the last appearance datum of Sphaeroidinellop molluscs (Bathytoma cataphracta, Calcarata calcara

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Fig. 3. GAMPS-00924, lateral tooth of Pachyscyllium distans (Probst, 1879) from Certaldo (Tuscany, central Italy). A – Lingual view; B – distal view; C – apical view; D – mesial view; E – labial view.

ta, Pelecyora brocchii, Petaloconchus intortus, Tena Pachyscyllium distans (Probst, 1879) godus obtusus, and Thylacodes arenarius, among oth- Figs. 3, 4 ers) and a single species of acorn barnacle (Concavus concavus). Vertebrate fossils from the Certaldo quarry Synonymy: See Reinecke et al. (2011: 52), where this spe- include the scyliorhinid specimen studied herein, as cies is referred to as Premontreia (Oxyscyllium) distans well as a few other teeth of elasmobranchs (assigned (Probst, 1879); for an updated list of synonyms and other to Megascyliorhinus miocaenicus and Rostroraja sp.) selected references, include also: and indeterminate otoliths. Most of these remains, in- ?1949 Scyliorhinus guttatum Probst, 1879. – Bauzá cluding the specimen described herein, lack a precise Rullán, p. 208, pl. 15, figs. 1, 2. stratigraphic position; they rolled down from the eas- 1953 Scylliorhinus (= Scyllium) sp. – Bauzá Rullán, ily erodible cliff to accumulate at its base (Fig. 2B). p. 5 (in part), figs. 7–14. 1976 Scyliorhinus distans (Probst, 1879) – de Ceuster, p. 136, pl. 5, figs. 13, 14. ?1992 Scylliorhinus distans (Probst, 1879). – Marín, p. 12, pl. 1, figs. 6, 7. 3. Systematic palaeontology 1996 Scyliorhinus distans Probst, 1879. – Mora Morote, p. 109, pl. 7, figs. 1–2, 4, 5. Class Chondrichthyes Huxley, 1880 2009 Pachyscyllium dachiardi (Lawley, 1876) – Briss- walter, p. 32 (in part; personal observation by Subclass Elasmobranchii Bonaparte, 1838 F.H.M., Elasmobranch Research Belgium collec- Order Carcharhiniformes Compagno, 1973 tions – Cabrières d’Aigues), table 3a. Family Scyliorhinidae Gill, 1862 2011 Pachyscyllium aff. dachiardii (Lawley, 1876). – Subfamily Premontreinae Cappetta, 1992 Vialle et al., p. 250 (in part), figs. ?3.8, 3.9. Genus Pachyscyllium Reinecke, Moths, Grant & 2012 Premontreia distans (Probst, 1879). – Bor et al., p. 41, pl. 22. Breitkreutz, 2005 2014 Pachyscyllium distans (Probst, 1879). – Reinecke, p. 7. Type species: Pachyscyllium albigensis Reinecke, Moths, 2019 Pachyscyllium distans (Probst, 1879). – Everaert Grant & Breitkreutz, 2005. et al., table 1, pl. 3, figs. 1, 2.

Fig. 4. GAMPS-00924, lateral tooth of Pachyscyllium distans (Probst, 1879) from Certaldo (Tuscany, central Italy). A – General view of the labial aspect of the tooth; B–D – four details of the labial aspect of the crown, with a focus on the longitudinal labial ridges that are found at the crown base. Note that the longitudinal labial ridges ornamenting the lateral cusplets (panels C and D) are more salient than those located at the base of the main cusp (panel B). Scanning Electron Mi- croscope (SEM) images by Karen Gariboldi, taken at the Dipartimento di Scienze della Terra dell’Università di Pisa (Pisa, Italy) using a Hitachi TM3030Plus Tabletop Microscope operating in the back-scattered electron mode.

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Taxonomic caveat: Pachyscyllium distans has a some- Table 1. Measurements of GAMPS-00924, lateral tooth of what troubled taxonomic history. As several other extinct Pachyscyllium distans (Probst, 1879) from the Pliocene of scyliorhinids, it was originally described by Probst (1879) Certaldo (Tuscany, central Italy). All measurements are re- as a member of Scyllium; the latter is a junior synonym of ported in millimetres. the living genus Scyliorhinus, to which Scyllium distans was transferred by Cappetta et al. (1967). Later, Cappetta GAMPS-00924 (2006; see also preliminary comments in Cappetta 1992) Apicobasal tooth height 2.8 reassigned Scyliorhinus distans to the extinct genus and sub- Mediolateral tooth width 3.2 genus Premontreia (Oxyscyllium), hence the new combina- Labiolingual tooth thickness 1.7 tion Premontreia (Oxyscyllium) distans. Finally, Reinecke (2014) transferred Probst’s species to the fossil premontreine genus Pachyscyllium, a view that is followed here. Measurements: Measurements of GAMPS-00924 are reported in Table 1. Referred material: One isolated tooth, currently kept at Ba- dia a Settimo (Scandicci, Italy) in the permanent exhibition Remarks on the species- level attribution: Some authors of “Gruppo AVIS Mineralogia e Paleontologia Scandicci” (e.g., Cappetta 2006; Cappetta 2012) consider P. distans (GAMPS) under accession number GAMPS-00924. as a junior synonym of Pachyscyllium dachiardii (= Scylli Occurrence: Pliocene (3.98–3.19 Ma) open shelf mud- um D’Achiardii) – a view first proposed byLandini (1977). stones exposed at Certaldo, Florence Province, Tuscany, Scyllium D’Achiardii was erected by Lawley (1876) on the central Italy. basis of eleven fossil teeth from the Pliocene deposits of Tuscany; unfortunately, the original description of this ex- Description: The crown of this tooth has a main central cusp tinct species was not accompanied by any figure depicting flanked up by a pair of lateral cusplets. In labial view, the the type series. Lawley (1876: 36) described the teeth of stout main cusp is distinctly triangular and moderately pro- P. dachiardii (often misspelled as “dachiardi” in later publi- jects distally; in lateral view, it exhibits a weakly sigmoidal cations) as follows: profile as its tip moderately bends labially. The labial surface “Alcuni denti tricuspidati, privi di qualunque seghettatu- of the main cusp is slightly convex transversely, although its ra, molto rigonfi alla base e che vanno terminando in pun- limit appears as concave in occlusal view; in turn, the lin- ta, la quale si rivolta dalla parte interna, portanti un piccolo gual face is invariantly strongly convex. Whereas the lingual dentino per lato similmente rigonfio e appuntato senza trac- surface of the main cusp is substantially smooth, the basal cia in qualunque parte della base di solchi longitudinali, con part of the labial surface displays obvious vertical ridges; the radice bassa e piana […].” latter are closely spaced and parallel to each other. The labial A translation of Lawley’s text is here proposed: base of the main cusp moderately overhangs the labial face “Some tricuspidate teeth, devoid of any serrations, strong- of the root. Both the mesial and the distal cutting edges of the ly bulging at their basis and tapering towards their apexes, main cusp are unserrated and complete; the mesial cutting which are buccally recurved; [teeth] bearing a similarly edge is gently convex, whereas the distal cutting edge is very bulging and pointed small denticle at each side, without any weakly concave. The lateral cusplets are continuous with trace of longitudinal sulci on any part of the basis, with low (although well-set apart from) the main cusp; they are ro- and flat root […].” bust, relatively high, and similar to each other, being shaped Unfortunately, only one out of the eleven Lawley’s as onion domes. The lateral cusplets weakly diverge from specimens of P. dachiardii is currently known from the each other and the main cusp by projecting respectively api- palaeontological collections of the Museo di Storia Natu- comedially (the mesial cusplet) and apicodistally (the distal rale dell’Università di Pisa (Landini 1977). This specimen, cusplet). Both the mesial and the distal cutting edges of the which was first figured by Landini (1977), conforms well main cusp prosecute through the lateral cusps to reach the to the description provided by Lawley (1876) and, crucial- tooth root, thus forming a sharp, continuous, mediolateral ly, exhibits a smooth labiobasal margin of the crown. By crown contour. Strong, salient, carina-like longitudinal ridg- reading Lawley’s original description, it is clear that all es, distinctly stronger than those observed at the labial base the eleven teeth comprising the type series of P. dachiardii of the main cusp, mark the labial and labiolateral surfaces of showed the same character, which Lawley (1876) regarded both the lateral cusplets. The root is rather robust and holau- as crucial for distinguishing this species from Scyllium pau lacorhize; it is comprised of two lobes that flare laterobasally luccii (a junior synonym of the living Mediterranean species and are separated from each other by a relatively narrow but Scyliorhinus stellaris according to Bassani 1901). In turn, well- marked furrow. The basal surface of the root is remark- one of the most striking features of the sole diagnostic syn- ably flat and somewhat heart- shaped. On the lingual face of type of Pachyscyllium distans (Reinecke et al. 2011: text- the root, a deep, longitudinal, nutrient groove bisects a rather fig. 17) relies in the presence of well- distinct longitudinal prominent, horizontally elongated lingual bulge. labial ridges at the crown base. Personal observations by the GAMPS-00924 is recognised as a lateral tooth based on present authors on an unpublished tooth set of Pachyscylli the observation of a rather stout crown whose main cusp um from some Pliocene localities of Tuscany (including the is distinctly bent distally (Reinecke et al. 2011; Cappetta lower Pliocene vertebrate-bearing site of Arcille; Bianucci 2012). According to Reinecke et al. (2011) and Bor et al. et al. 2019) revealed that the presence of a smooth labiobas- (2012), the observation of a gently convex (rather than al margin of the crown represents a stable dental character straight) mesial cutting edge might be related to a low de- of Lawley’s species, GAMPS-00924 being indeed the sole gree of dignathic heterodonty.

eschweizerbart_xxx Pachyscyllium distans 135 examined Tuscan Pliocene Pachyscyllium tooth that bears of P. distans from the Mediterranean Pliocene, all of “distans- like” longitudinal labial ridges at the crown base. which are from Spanish localities and were published Consequently, we agree with Reinecke et al. (2011) in re- on magazines of natural sciences that are not widely garding P. distans as a valid, separate species that significantly differs from P. dachiardii (and not just a result of distributed outside Spain. The first of this records was sexual dimorphism or any other form of intraspecific het- provided by Bauzá Rullán (1949), who reported erodonty). on one tooth identified as belonging to Scyliorhinus guttatum (currently recognised as a junior synonym of Chaenogaleus affinis; Reinecke et al. 2011) from supposedly Pliocene deposits of Son Talapí, Mallor- 4. General discussion and conclusions ca, Balearic Islands. This tooth seemingly displays a typically premontreine outline (Bauzá Rullán 1949: A scrutiny of the palaeoichthyological literature re- figs. 1, 2) and was described as exhibiting small verti- veals that teeth of Pachyscyllium distans have been cal folds at the crown base; as such, it likely represents reported from several Neogene sites of the Mediter- a specimen of P. distans. That said, the stratigraphic ranean, Paratethys, and the Northeastern Atlantic whereabouts of this find are somewhat uncertain, as (palaeo-North Sea and Portugal). Pachyscyllium dis demonstrated by some inconsistencies in later works tans has also been recorded from lowermost Miocene by the same author [e.g., Bauzá Rullán 1955, in (Aquitanian) deposits of the Northwestern Atlantic which both the terms “Plaisanciense” (= Piacen- (Eastern coast of the USA) (see Case 1980), but these zian) and “Plioceno inferior” (= lower Pliocene, cor- strata (i.e., the Trent Marl Formation at River Bend responding to the Zanclean stage) are applied to the Plantation, Craven County), are here considered to be Scyliorhinus guttatum locality]. A second possible of Oligocene age (see Ward 1978), and thus not dealt record of P. distans from the Mediterranean Pliocene with in this study. The same applies for other Oligo- was provided by Marín (1992), who reported on teeth cene records of Pachyscyllium aff. distans from Eu- of Scylliorhinus [sic] distans from presumably lower rope (see e.g. Reinecke et al. 2014). Pliocene (Zanclean) deposits of Butano, near Elche, With respect to the Mediterranean and Paratethyan Alicante Province. Marín (1992) did not describe regions, Pachyscyllium distans has frequently been these teeth, and the poor quality of his illustrations collected from Miocene strata. In particular, as regards (Marín 1992: pl. 1, figs. 6, 7) does not allow a positive the Paratethyan sectors, P. distans has commonly been identification of these specimens as belonging either reported from the upper lower Miocene (i.e., Burdiga- to P. distans or to P. dachiardii, which in turn is well- lian) of southern Germany, Switzerland, and Austria, known from various Mediterranean Pliocene locali- as well as from the middle Miocene of Austria and ties (Lawley 1876; Bauzá Rullán 1953: figs. 1–6; Poland, whereas the Mediterranean records are mostly Landini 1977; Bellocchio et al. 1991; Mañe et al. from deposits of Aquitanian to Langhian age of France 1996; Cappetta & Cavallo 2006; Marsili 2008). Fi- [Reinecke et al. 2011, and references therein, but nally, Mora Morote (1996) described and illustrat- see Bauzá Rullán (1953) for a reliable occurrence ed teeth of Scyliorhinus distans from the Pliocene of of P. distans from supposedly lower upper Miocene Guardamar del Segura, near Alicante, southern Spain. (i.e., Tortonian) strata of Mallorca, Balearic Islands, The dental characters of these specimens, as reported Spain]. Under various names [i.e., Scyliorhinus dis by Mora Morote (1996), compare favourably with tans, Pachyscyllium dachiardi (error pro dachiardii), those of Pachyscyllium distans, and include allegedly and Pachyscyllium aff. dachiardii], Cappetta (1970: diagnostic traits such as the presence of longitudinal e.g. pl. 9, fig. 4),Brisswalter (2009), and Vialle et al. labial ridges at the crown base. As regards the prov- (2011: e.g. fig. 3.9) described and figured material of enance of these teeth, Adnet et al. (2010) reported P. distans from the middle Miocene of the Montpellier that the material studied by Mora Morote (1996) region, the Luberon Massif, and Mazan, respectively; comes from the Rojales Sandstone Formation, whose overall, these records from southeastern France testify geologically youngest marine strata (belonging to the to the presence of P. distans in the southern Rhodani- P2 system of Soria et al. 2005) bear intercalations of an Basin, a narrow seaway that might have connected palustrine marls that host rodent fossils dating into the the Mediterranean Sea to the western Paratethys (e.g., Mammal Neogene 14 zone; the latter has recently been Reynaud & James 2012). To our knowledge, besides calibrated by Minwer- Barakat et al. (2012) at the the present study, there are only three possible records ~5.3–4.19 Ma time span. Therefore, considering also

eschweizerbart_xxx 136 A. Collareta et al. the uncertainties associated with the taxonomic identi- of thermophilic marine vertebrates (e.g., Sorbi et al. fication and the geological age of the specimens men- 2012; Prista et al. 2013; Collareta et al. 2017b). tioned by Bauzá Rullán (1949) and Marín (1992), However, other events, such as the disruption of the GAMPS-00924 represents the second bona fide re- psychrospheric ecosystems (and related deep- water cord of P. distans from the Pliocene of the Mediterra- trophic webs) of the Mediterranean Sea (Cigala Ful- nean Sea. Therefore, our discovery contributes to our gosi 1996; Marsili 2007; Cigala Fulgosi et al. 2009; knowledge on the late Neogene central Mediterranean Collareta et al. 2019) might also explain the eventual biodiversity and encourages to pursue the quest for extirpation of P. distans and other elasmobranch taxa fossil elasmobranchs over the entire Tuscan Pliocene from this satellite basin of the North Atlantic. Hopeful- marine succession. ly, the collection of new specimens of P. distans from Outside the Mediterranean/Paratethyan realm, chronostratigraphically framed localities of the Tuscan P. distans has been recorded from various upper lower Pliocene will contribute to further shed light on this Miocene (Burdigalian) to ?upper Miocene (?Tortonian) issue and, more generally, on the palaeoecological and localities of Portugal (Antunes & Jonet 1970; Jonet palaeobiogeographic dynamics of the late Neogene 1978; Antunes et al. 1981). Pachyscyllium distans is elasmobranch faunas of the Mediterranean Sea. also known as a common component of the Miocene elasmobranch assemblages of the North Sea Basin, where it ranges chronostratigraphically between the Aquitanian and the Langhian (Reinecke et al. 2011; Acknowledgements Bor et al. 2012; Everaert et al. 2019, and references therein). Teeth of P. distans reported by de Ceuster We are very grateful to Mark Bosselaers, Bert Gijsen, (1976) from basal Pliocene deposits exposed at Rumst Kristiaan Hoedemakers, Jürgen Pollerspöck, and (near Antwerp, Belgium) are most likely reworked Thomas Reinecke, who provided precious comparative material and bibliographic support. Thomas Reinecke is from the lower to middle Miocene strata of the Ber- also warmly thanked for fruitful discussions over the is- chem Formation, as most of the fossil material from sues dealt with in the present paper. Thanks are also due that horizon (Reinecke et al. 2011; personal observa- to Karen Gariboldi, for having led the SEM analysis of tion by F.H.M.). In the light of these distributional data, GAMPS-00924, and to Giulia Bosio, for her field support. and considering also the aforementioned age estimates Last but not least, we are sincerely indebted to Giorgio Car- nevale, an anonymous reviewer, and Günter Schweigert, regarding the P. distans specimens described by Mora whose thorough reviews greatly contributed to improve an Morote (1996), GAMPS-00924 represents the geo- early draft of this paper. logically youngest confirmed occurrence of P. distans worldwide. Furthermore, it appears that P. distans was extirpated from the North Sea Basin at the end of the Langhian, whereas it persisted along the coasts of Por- References tugal till the Serravallian or Tortonian, and eventually survived into the Pliocene in the Mediterranean Sea Adnet, S., Balbino, A. C., Antunes, M.T. & Marín- only. This southward trend of Neogene regional ex- Ferrer, J.M. (2010): New fossil teeth of the White tinctions might suggest that, somewhat similar to what Shark (Carcharodon carcharias) from the Early Plio- has been hypothesised for the congeneric and largely cene of Spain. Implication for its paleoecology in the Mediterranean. – Neues Jahrbuch für Geologie und sympatric species P. dachiardii (Reinecke et al. 2011), Paläontologie, Abhandlungen, 256 (1): 7–16. doi: P. distans was a thermophilic catshark, whose range 10.1127/0077-7749/2009/0029 contracted southward as colder conditions progres- Antunes, M.T. & Jonet, S. (1970): Requins de lʼHelvétien sively took hold in the Northeastern Atlantic by effect supérieur et du Tortonien de Lisbonne. – Revista da Fac- of a progressively cooling global climate and follow- uldade de Ciências da Universidade de Lisboa, C–Ciên- cias naturais, (2), 16 (1): 119–280. ing the onset of the Northern Hemisphere glaciation Antunes, M.T., Jonet, S. & Nascimiento, A. (1981): (e.g., Herbert et al. 2016). If this interpretation is Vertébrés (crocodiliens, poissons) du Miocène marin correct, it seems reasonable to hypothesise that P. dis de lʼAlgarve occidentale. – Ciências da Terra (UNL), 6: tans went definitively extinct around 3 Ma, when the 9–38 Mediterranean region experienced a pulse of severe Bassani, F. (1901): Su alcuni avanzi di pesci del Pliocene to- scano. – Monitore Zoologico Italiano, 12 (7): 189–191. climate degradation (Prista et al. 2015, and references Bauzá Rullán, J. (1949): Contribuciones al conocimien- therein) that likely led to the extirpation of several taxa to de la fauna ictiológica fósil de Mallorca. – Boletin

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